Correlated patterns of variation in phenology and seed production in populations of two annual grasses along an aridity gradient

I applied a comparative approach to reveal correlated patterns of variation in phenology and seed production in four populations of two annual grasses Hordeum spontaneum and Avena sterilis, sampled in the same environments distributed along an aridity gradient in Israel. The steep aridity gradient in Israel represents two parallel clines of environmental productivity (annual rainfall) and predictability (variation in amount and timing of annual rainfall) that is likely to induce similar responses in natural plant populations distributed along the gradient, if (1) selection is strong, (2) species share the same ecological niche, and (3) there is genetic variation for ecologically important traits. I found in plants of both species (1) ultimate advance in onset of flowering, and (2) more but smaller seeds, with increasing aridity. The broad sense heritabilities of onset of flowering, seed size and seed yield in both species were very high, moderate and low, respectively. It appears that the observed adaptive complex of traits have evolved in both species in response to this specific array of environments.     

Community‐level consequences of species interactions in an annual plant community

Question: How does the intensity of species interactions affect species and functional group composition of an annual plant community? Location: Sede Boqer, Negev Desert, Israel. Methods: The potential for competitive interactions in two annual plant communities (desert and coastal) from semi‐stabilized sand dunes was manipulated by varying seed bank density and therefore the number of potentially interacting individuals. Communities were exposed to three different irrigation regimes, mimicking precipitation at the desert site, the coastal site, and an intermediate precipitation level. Plots were maintained for 3 years, and percentage cover of each species in the plots was recorded at the end of each growing season. We used redundancy analysis to test for effects of initial density, irrigation, and year on the species and functional group composition of the communities. Results: Initial density had significant effects on species composition, and these effects remained significant over 3 years, even as total community percentage cover became more similar among treatments over time. Density effects did not depend on resource availability (irrigation level). Functional group identity or individual plant size did not predict which species would be good competitors, and a species' competitive ability did not predict its abundance in the field. Conclusions: Species interactions strongly affect community composition, and those effects carry over into subsequent years such that competition does not lead to convergence in community structure over time. However, the particular changes in composition observed were not predictable by some of the traits that have been found important in individual‐level experiments. We speculate that the outcome of competition in diverse communities will depend on multiple traits, in contrast to the outcome of individual‐level pairwise experiments. We also speculate that the shift in composition with density could mean that local variation in density may contribute to maintenance of diversity in this system.     

Annual plant–shrub interactions along an aridity gradient

Annual plants in semi-arid and arid areas are often closely associated with shrubs. The degree of association largely depends on the balance of negative and positive effects between these contrasting plant life-forms, ranging from interference to facilitation. Since positive interactions are predicted to become less important with increasing rainfall, the interaction balance is expected to shift along aridity gradients. However, this prediction has not been tested on a community level and for different life-history stages across large geographical gradients. Here, we show such changes for annual plant populations and communities in four contrasting sites along a steep climatic gradient, ranging from the arid desert to mesic Mediterranean regions in Israel. Above-ground productivity, richness, seedling density, and seed bank density of the annual plant community, as well as fecundity of annual plant populations, were generally higher under shrubs than in areas between shrubs at the arid end of the gradient, but significantly lower at the humid end. Net effects of shrubs on annuals expressed as relative interaction intensity indicated a steady and consistent shift from net positive or neutral effects in the desert to net negative effects in the mesic part of the gradient. These findings emphasize the usefulness of studies along large-scale gradients encompassing a wide range of environmental conditions for understanding community level interactions among coexisting species.     

Ecotypic differentiation in Medicago polymorpha L. along an environmental gradient in central Chile. I. Phenology,biomass production and reproductive patterns

Burr Medic (Medicago polymorpha) is awidespread, polymorphic annual legume of Mediterranean origin, and as aself-reseeding, highly effective nitrogen fixer it has important value invarious dryland farming systems. In central Chile, it is naturalized along a1000 km long environmental gradient, from an arid zone(29°–31° S) to a perhumid one(37°–38°24' S). Comparative and experimental studiesof its ecotypic differentiation patterns along this gradient should be usefulinelucidating the evolutionary forces at work along environmental gradients, andalso for efforts to select annual legumes for pastures and fallow fieldimprovement in mediterranean climate areas. In this first paper of atwo-part series, we present results of two common-environmentexperiments conducted at a subhumid site in central Chile to compare vegetativeand reproductive traits in 69 populations of Burr Medic collected in 1988 and1994. Multivariate analyses based on 11 phenological and agronomic characters,including phenology and biomass accumulation, winter vigour and degree ofhard-seededness, showed that three principal components (PCs)explained nearly 70% of total variability, in both collections. For thelarger collection (1988), the diagram of dispersion for the first twoPCs – phenology, winter vigour and above-ground biomass –provided good discrimination between accessions from arid-semiarid andhumid-perhumid mediterranean zones. Onset of flowering was positivelycorrelated with both latitude and longitude, as well as mean annualprecipitation/potential evapotranspiration (PP/ETP) ofcollecting site; days from first flower to pod ripening was negativelycorrelated with PP/ETP. In contrast to reports from Syria and Sardinia, noclear trends in Burr Medic seed or pod-related traits was found along theChilean gradient, apart from the concentration of spiny pod accessions in thesouthern, more mesic end of the gradient. Winter vigour (according to asemi-quantitative analysis of growth) was greater in accessions fromthe warmer, arid and semiarid zones than in those from cooler, subhumid tohumidones. Both winter vigour and harvest index were negatively correlated with daysto first flower. Applied and evolutionary aspects of the results are brieflydiscussed.     

Seed mass and dormancy of annual plant populations and communities decreases with aridity and rainfall predictability

Several theoretical and empirical studies have examined the influence of environmental conditions on seed traits and germination strategies of annual species. A positive relationship between seed mass and dormancy has been described for annuals occupying climatically unpredictable ecosystems. Larger-seeded species tend to have higher seedling survival rates, while dormancy allows a bet-hedging strategy in unpredictable environments. Until now, these ideas have been addressed primarily for only one or a few focal species, without considering differences among populations and communities. The novelty of the present study lies in the population and community-level approach, where a comprehensive seed trait database including 158 annual species occurring along a gradient of rainfall variability and aridity in Israel was used to ask the following question: Does average seed mass and dormancy of annual populations and communities decrease with increasing aridity and rainfall unpredictability?Soil seed bank samples were collected at the end of the summer drought, before the onset of the rains, from four plant communities. Germination was tested under irrigated conditions during three consecutive germination seasons to determine the overall seed germinability in each soil sample. Seed mass was obtained from newly produced seeds collected at the study sites in late spring. The community level results showed that, in contrast to common theoretical knowledge, seed mass and dormancy of the dominant annual species decreased with increasing aridity and rainfall variability. Accordingly, a negative correlation was found between seed mass and seed germination fractions. The present study demonstrates that an analysis of seed traits along climatic gradients is significantly improved by approaches that target both population and community levels simultaneously. A critical evaluation sheds new light upon the selective pressures that act on seed ecology of annuals along a climatic gradient and facilitates formulation of more mechanistic hypotheses about factors governing critical seed traits.     

Reproductive allocation strategies in desert and Mediterranean populations of annual plants grown with and without water stress

Reproductive effort (relative allocation of biomass to diaspore production) was compared in matched pairs of Mediterranean and desert populations of three unrelated annual species, Erucaria hispanica (L.) Druce, Bromus fasciculatus C. Presl. and Brachypodium distachyon (L.) Beauv., grown under high and low levels of water availability in a common-environment experiment. Desert populations in all three species showed higher reproductive effort than corresponding Mediterranean populations, as expressed by both a reproductive index (RI= reproductive biomass/vegetative biomass), and a reproductive efficiency index (REI=number of diaspores/total plant biomass). Moreover, in E. hispanica and Brachypodium distachyon, inter-populational differences in reproductive effort were greater under water stress, the main limiting factor for plant growth in the desert. These results indicate that variability in reproductive effort in response to drought is a critical and dynamic component of life history strategies in annual species in heterogeneous, unpredictable xeric environments. When subjected to water stress the Mediterranean populations of E. hispanica and B. distachyon showed greater plasticity (e.g. had a greater reduction) in reproductive effort than the desert populations, while in Bromus fasciculatus both populations showed similar amounts of plasticity.     

Germination strategies under climate change scenarios along an aridity gradient

由于年降雨量减少和变异性的增加,地中海东部地区的气候变化将对生态系统功能和植物群落动态产生重大影响。我们旨在了解种子库作为应对气候变化所导致的气候不确定性的潜在缓冲作用。我们研究了沿干旱梯度出现的18种常见物种的萌发策略。数据由干旱、 半干旱、地中海和中等地中海生态系统连续九年内萌发的土壤种子库获得。在半干旱和地中海地区,采用了模拟30%干旱和30%降雨增加的降雨处理方法。在连续三个萌发季的最佳灌溉条件下检测了萌发策略,以确定每种土壤样品的总体种子萌发能力。使用一种新颖的统计方法研究了萌发策略的变化,该方法考虑了可能影响种子发芽性的气候和生物因素。研究结果表明,优势种通过产生具有不同年度发芽率概率的种子来控制其发芽率。可产生种子的降雨量导致了关于可萌发性的两种主要种子类型：高萌发性(可形成短暂种子库的种子)和低萌发性(可形成持久种子库的种子)。我们得出的结论是,两种类型的干湿年之间种子产生的差异沿干旱梯度建立了一个稳定的平衡,使土壤种子库可以充当稳定机制,以防止降雨的不可预测性。此外,我们提出了地中海和干旱生态系统中占主导地位的一年生物种萌发策略的一般模型,该模型加强了土壤种子库可以作为应对该地区气候变化引起的气候不确定性的缓冲剂的概念。     

Population differentiation in an annual legume: local adaptation

Abstract. Studies of many plants species have demonstrated adaptive genetic differentiation to local environmental conditions. Typically these studies are conducted to evaluate adaptation to contrasting environments. As a consequence, although local adaptation has been frequently demonstrated, we have little information as to the spatial scale of adaptive evolution. We evaluated adaptive differentiation between populations of the annual legume Chamaecrista fasciculata using a replicated common-garden design. Study sites were established in three field locations that are home to native populations of C. fasciculata . Each location was planted for two years with seed from the population native to the study site (home population) and populations located six distances (0.1-2000 km) from each site (transplanted populations). Seeds were planted into the study sites with minimum disturbance to determine the scale of local adaptation, as measured by a home-site fitness advantage, for five fitness components: germination, survival, vegetative biomass, fruit production, and the number of fruit produced per seed planted (an estimate of cumulative fitness). For all characters there was little evidence for local adaptation, except at the furthest spatial scales. Patterns of adaptive differentiation were fairly consistent in two of the three sites, but varied between years. Little genetic variation was expressed at the third site. These results, combined with previous estimates of limited gene flow, suggest that metapopulation processes and temporal environmental variation act together to reduce local adaptation, except over long distances.     

Ecotypic differentiation in Medicago polymorpha L. along an environmental gradient in central Chile. II. Winter growth as related to phenology and temperature regime

In this second article of a series analysing ecotypic differentiation ofthe Burr Medic (Medicago polymorpha L.),occurring along a 1000 km natural aridity gradient in centralChile, we report variations in growth potential at low temperature in relationto phenology and latitude of collecting site. In two experiments conducted inoutdoor nursey trials at Cauquenes, in the 'intermediate'(i.e., subhumid) mediterranean zone (35°58 S,72°17 W), biomass accumulation among 4 and 18 accessions ofBurrMedic, was compared. In Experiment 1, the two early-flowering accessionscollected in the arid and semiarid zones (northern Chile) showedsignificantly greater winter growth potential than the twolate-floweringaccessions from humid, south-central Chile, at five different sowingdensities. In Experiment 2, winter growth was compared, at a single sowingdensity, among 18 accessions collected in various sites along the ariditygradient. Here again, growth patterns were closely correlated to latitude ofcollecting site, with early-flowering ecotypes from the north showingsignificantly greater winter growth potential than those from the south. In athird experiment, relative growth rate, net assimilation rate and leaf arearatio and phytomass production were compared among eight accessions grown undertemperature regimes simulating natural growth conditions at Cauquenes in June(winter) and October (spring), i.e.,12°/7°Cand 18°/13°C day/night, respectively. Asbefore, northern accessions showed higher growth potential in the lowertemperature regime than those from humid and perhumid areas at more southernlatitudes. These results suggest that in arid to semiarid environments, whererainfall is erratic and unpredictable, ecotypes of Burr Medic are naturallyselected which show early-flowering and high growth capacity duringwinter, the period of the year when rainfall is most likely to occur inmediterranean-climate regions. Conversely, in humid areas, with higherrainfall predictability, and higher frequency of killing frost, ecotypes havebeen selected with longer periods of growth potential but lower winter growth.Applied and ecological aspects of the results are discussed in the context of along-term program of annual pasture improvement programme.     

Inter-population variability of leaf morpho-anatomical and terpenoid patterns of Pistacia atlantica Desf. ssp. atlantica growing along an aridity gradient in Algeria

Three Algerian populations of female Pistacia atlantica shrubs were investigated in order to check whether their terpenoid contents and morpho-anatomical parameters may characterize the infraspecific variability. The populations were sampled along a gradient of increasing aridity from the Atlas mountains into the northwestern Central Sahara.As evidenced by Scanning Electron Microscopy, tufted hairs could be found only on seedling leaves from the low aridity site as a population-specific trait preserved also in culture. Under common garden cultivation seedlings of the high aridity site showed a three times higher density of glandular trichomes compared to the low aridity site. Increased aridity resulted also in reduction of leaf sizes while their thickness increased. Palisade parenchyma thickness also increases with aridity, being the best variable that discriminates the three populations of P. atlantica.Analysis of terpenoids from the leaves carried out by GC-MS reveals the presence of 65 compounds. The major compounds identified were spathulenol (23 μg g⁻¹ dw), α-pinene (10 μg g⁻¹ dw), verbenone (7 μg g⁻¹ dw) and β-pinene (6 μg g⁻¹ dw) in leaves from the low aridity site; spathulenol (73 μg g⁻¹ dw), α-pinene (25 μg g⁻¹ dw), β-pinene (18 μg g⁻¹ dw) and γ-amorphene (16 μg g⁻¹ dw) in those from medium aridity and spathulenol (114 μg g⁻¹ dw), α-pinene (49 μg g⁻¹ dw), germacrene D (29 μg g⁻¹ dw) and camphene (23 μg g⁻¹ dw) in leaves from the high aridity site. Terpene concentrations increased with the degree of aridity: the highest mean concentration of monoterpenes (136 μg g⁻¹ dw), sesquiterpenes (290 μg g⁻¹ dw) and total terpenes (427 μg g⁻¹ dw) were observed in the highest arid site and are, respectively, 3-, 5- and 4-fold higher compared to the lower arid site. Spathulenol and α-pinene can be taken as chemical markers of aridity. Drought discriminating compounds in low, but detectable concentrations are δ-cadinene and β-copaene. The functional roles of the terpenoids found in P. atlantica leaves and principles of their biosynthesis are discussed with emphasis on the mechanisms of plant resistance to drought conditions.     

Phenology and polyploidy in annual Brachypodium species (Poaceae) along the aridity gradient in Israel

Local adaptation of plants along environmental gradients provides strong evidence for clinal evolution mediated by natural selection. Plants have developed diverse strategies to mitigate stress, for example, drought escape is a phenological strategy to avoid drought stress, while polyploidy was proposed as a genomic adaptation to stress. Polyploidy as an adaptation to aridity (an environmental parameter integrating temperature and precipitation) was previously documented in annual Brachypodium spp. (Poaceae) in the Western Mediterranean. Here, we examined whether polyploidy or phenology are associated with aridity in annual Brachypodium spp. along the aridity gradient in the Eastern Mediterranean. Using flow cytometry, we determined ploidy levels of plants from natural populations along the Israeli gradient, spanning ∼424 km from mesic Mediterranean to extreme desert climates. In a common garden we recorded time of seedling emergence, flowering and senescence. We tested whether the proportion of allotetraploids in the populations and phenological traits were associated with aridity. Contrary to a previous study in the Western Mediterranean, we found no effect of aridity on the proportion of allotetraploids and diploids within populations. Interestingly, phenology was associated with aridity: time of emergence was later, while flowering and senescence were earlier in desert plants. Our results indicate that in the Eastern Mediterranean, adaptation of Brachypodium to aridity is mediated mainly by phenology, rather than ploidy level. Therefore, we suggest that genome duplication is not the main driver of adaptation to environmental stress; rather, phenological change as a drought escape mechanism may be the major adaptation.     

Life history variation in an annual plant under two opposing environmental constraints along an aridity gradient

Environmental gradients represent an ideal framework for studying adaptive variation in the life history of plant species. However, on very steep gradients, largely contrasting conditions at the two gradient ends often limit the distribution of the same species across the whole range of environmental conditions. Here, we study phenotypic variation in a winter annual crucifer Biscutella didyma persisting along a steep gradient of increasing rainfall in Israel. In particular, we explored whether the life history at the arid end of the gradient indicates adaptations to drought and unpredictable conditions, while adaptations to the highly competitive environment prevail at the mesic Mediterranean end. We examined several morphological and reproductive traits in four natural populations and in populations cultivated in standard common environment. Plants from arid environments were faster in phenological development, more branched in architecture and tended to maximize reproduction, while the Mediterranean plants invested mainly in vertical vegetative growth. Differences between cultivation and field in diaspore production were very large for arid populations as opposed to Mediterranean ones, indicating a larger potential to increase reproduction under favorable conditions. Our overall findings indicate two strongly opposing selective forces at the two extremes of the aridity gradient, which result in contrasting strategies within the studied annual plant species.     

Phenological and morphological differentiation in annual Chaetanthera moenchioides (Asteraceae) over an aridity gradient

Flowering-time, plant longevity and size of capitulum were studied in a common garden experiment on seven populations of annual Chaetanthera moenchioides derived from a strong latitudinal aridity gradient in the mediterranean climate area of Chile. Populations derived from the drier northern part of the latitudinal gradient showed significantly shorter flowering and fruiting phenology and smaller capitula under uniform growing conditions. Water stress experiments performed in the greenhouse induced further phenological hastening and a reduction in number of ray florets per capitulum. Population differentiation and the reaction norms in C. moenchioides agree with the predictions of genetic-assimilation given that the plastic response of the species under water stress mimics phenotypic differentiation that has evolved along the environmental gradient.     

Variation in onset of summer dormancy and flowering capacity along an aridity gradient in Poa bulbosa L., a geophytic perennial grass

Variation in the onset of summer dormancy and flowering capacity of 16 populations of Poa bulbosa, collected along a steep north-south aridity gradient in Israel (810-110 mm rain year(-1)), was studied under controlled conditions in a phytotron (16 h daylength, 22/16 degrees C day/night) and under natural conditions in a garden experiment in a net-house. Plant age at the onset of dormancy varied markedly amongst populations (7-16 weeks under controlled conditions) and was positively correlated with mean annual precipitation at the site of origin of the population, i.e. dormancy was earlier as aridity increased. Flowering capacity in the different populations was negatively correlated with rainfall in the original habitat and, consequently, also with the age at onset of dormancy, i.e. the lower the mean annual precipitation, the earlier the onset of dormancy and the higher the proportion of flowering plants and panicles per plant. Differences in xeromorphic leaf traits were also observed among populations from locations differing in aridity. Plants from the more arid sites (110-310 mm year(-1)) generally had greyish and curved leaves, whereas plants from more humid sites (500-810 mm year(-1)) tended to have green and straight leaves. Thus, plants with curved and/or greyish leaves generally had a higher flowering capacity and entered dormancy earlier than plants with straight and/or green leaves. The significance of the association among these traits for the adaptation of P. bulbosa to increasing aridity is discussed.     

Nuclear and cytoplasmic contributions to intraspecific divergence in an annual legume

The genetic architecture of trait differentiation was evaluated between two ecologically distinct populations of Chamaecrista fasciculata. Individuals from Maryland and Illinois populations were crossed to create 10 types of seed: Maryland and Illinois parents, reciprocal F1 and F2 hybrids, and backcrosses to Maryland and to Illinois on reciprocal F1 hybrids. Reciprocal crosses created hybrid generation seeds with both Maryland and Illinois cytoplasmic backgrounds. Experimental individuals were grown in a common garden near the site of the Maryland population. In the garden, plants from the Illinois population flowered, set fruit, and died earlier than those from Maryland, likely reflecting adaptations to differences in growing season length between the two populations. Although reproductive components at the flower and whole plant level differed between the two populations, reproductive output as measured by fruit and seed production was similar. Cytoplasmic genes had a subtle but pervasive effect on population differentiation; hybrids with Maryland cytoplasm were significantly differentiated from those with Illinois cytoplasm when all characters were evaluated jointly. The nuclear genetic architecture of population differentiation was evaluated with joint scaling tests. Depending on the trait, both additive and nonadditive genetic effects contributed to population differentiation. Intraspecific genetic differentiation in this wild plant species appears to reflect a complex genetic architecture that includes the contribution of additive, dominance, and epistatic components in addition to subtle cytoplasmic effects.     

Repeated intraspecific divergence in life span and aging of African annual fishes along an aridity gradient

Life span and aging are substantially modified by natural selection. Across species, higher extrinsic (environmentally related) mortality (and hence shorter life expectancy) selects for the evolution of more rapid aging. However, among populations within species, high extrinsic mortality can lead to extended life span and slower aging as a consequence of condition‐dependent survival. Using within‐species contrasts of eight natural populations of Nothobranchius fishes in common garden experiments, we demonstrate that populations originating from dry regions (with short life expectancy) had shorter intrinsic life spans and a greater increase in mortality with age, more pronounced cellular and physiological deterioration (oxidative damage, tumor load), and a faster decline in fertility than populations from wetter regions. This parallel intraspecific divergence in life span and aging was not associated with divergence in early life history (rapid growth, maturation) or pace‐of‐life syndrome (high metabolic rates, active behavior). Variability across four study species suggests that a combination of different aging and life‐history traits conformed with or contradicted the predictions for each species. These findings demonstrate that variation in life span and functional decline among natural populations are linked, genetically underpinned, and can evolve relatively rapidly.     

Patterns and causes of spatial variation in the reproductive success of a desert annual

Summary This study examines patterns and causes of variation in the reproductive success of the desert annual Stipa capensis. Three nested scales of variation were analyzed: variation between individuals of the same plot, variation between different plots of the same habitat, and variation between different habitats in the same region. Perturbation experiments (irrigation and neighbors removal) were performed to test the effects of heterogeneity in soil water and neighborhood competition on the magnitude of variation in each scale. The results demonstrate that variation of reproductive success was highest within plots, lowest between plots, and moderate between habitats. Soil water heterogeneity contributed to spatial variation in all scales but was most important for differences between habitats. Neighborhood competition increased the variation within plots, but decreased the variation between habitats. The results further demonstrate that water limitation was negatively correlated with the position of the habitat along the run-off/run-on gradient. An opposite trend was obtained for the effect of competition.     

Soil property,CO2 emission and aridity index as agroecological indicators to assess the mineralization of cover crop green manure in a Mediterranean environment

In this study soil chemical and biochemical properties, cover crop biomass production and quality, and climatic factors (AI) have been taken into account in order to identify sensitive agroecological indicators suitable for an early assessment of green manuring outcomes, measured in terms of soil CO₂ emission and soil mineralization dynamics in a short term experiment in a Mediterranean environment. The field experiment was conducted over two cropping rotations during 2004–2005 in central Italy. A winter cover crop/sweet pepper sequence with the cover crop used as green manure was adopted. The cover crop treatments were common vetch (CV), rye grass (RG), and fallow soil as the control (Control). Soil enzyme activities (acid phosphatase. protease and β-glucosidase), CO₂ emission, and inorganic nitrogen concentrations were monitored from cover crop green manure incorporation to pepper harvesting in order to evaluate soil mineralization dynamics. The climatic conditions were summarized by the monthly aridity index (AI) calculated as the precipitation/temperature ratio. A group of mineralization indexes, calculated using values of available nitrogen and enzyme activities, was used to describe the soil process during crop cycle after green manure. The mineralization process dynamic results as a combined effect of climatic conditions and soil organic matter quality produced by different cover crop green manures. The common vetch green manuring was effective in lowering the soil C/N with respect to the control soil (5.7 vs. 8.3 and 8.5 vs. 12.1 in 2004 and 2005, respectively), promoting CO₂ emission (8.95 vs. 5.19 and 6.75 vs. 4.28 Mg CO₂-C ha⁻¹ in 2004 and 2005, respectively), enzyme activity, nitrogen release, and crop aboveground biomass (8.59 vs. 7.05 Mg ha⁻¹ dry matter). Among the selected agroecological indicators, the relationships between enzyme activities and the monthly aridity index may suggest a new approach for agronomists and soil scientists to understand the combined effect of temperature and precipitation on soil mineralization dynamic. The high aridity index at the time of green manuring may have caused a priming effect of SOM and promoted soil mineralization during the vegetable crop growing season in a Mediterranean environment. Finally, no evidence was found between soil CO₂ emission and the aridity index; soil respiration was mainly affected by cover crop biomass and the soil C/N ratio.